What are we protecting? Fisher behavior and the unintended consequences of spatial closures as a fishery management tool

Spatial closures like marine protected areas (MPAs) are prominent tools for ecosystem-based management in fisheries. However, the adaptive behavior of fishermen, the apex predator in the ecosystem, to MPAs may upset the balance of fishing impacts across species. While ecosystem-based management (EBM) emphasizes the protection of all species in the environment, the weakest stock often dominates management attention. We use data before and after the implementation of large spatial closures in a North Pacific trawl fishery to show how closures designed for red king crab protection spurred dramatic increases in Pacific halibut bycatch due to both direct displacement effects and indirect effects from adaptations in fishermen's targeting behavior. We identify aspects of the ecological and economic context of the fishery that contributed to these surprising behaviors, noting that many multispecies fisheries are likely to share these features. Our results highlight the need either to anticipate the behavioral adaptations of fishermen across multiple species in reserve design, a form of implementation error, or to design management systems that are robust to these adaptations. Failure to do so may yield patterns of fishing effort and mortality that undermine the broader objectives of multispecies management and potentially alter ecosystems in profound ways.     

Incorporation of Recreational Fishing Effort into Design of Marine Protected Areas

Abstract:  Theoretical models of marine protected areas (MPAs) that explore benefits to fisheries or biodiversity conservation often assume a dynamic pool of fishing effort. For instance, effort is homogenously distributed over areas from which subsets of reserves are chosen. I tested this and other model assumptions with a case study of the multiple-use Jervis Bay Marine Park. Prior to zoning of the park I conducted 166 surveys of the park's recreational fisheries, plotting the location of 16,009 anglers. I converted these plots into diagrams of fishing effort and analyzed correlates between fishing and habitat and the effect of two reserve designs—the draft and final zoning plans of the park—on the 15 fisheries observed. Fisheries were strongly correlated with particular habitats and had negatively skewed and often bimodal spatial distribution. The second mode of intensely fished habitat could be 6 SD greater than the fishery's mean allocation of effort by area. In the draft-zoning plan, sanctuary zone (no-take) area and potential subduction of fishing effort were similar. In the final plan, which was altered in response to public comment, the area of sanctuary zone increased, and the impact on fishing effort decreased. In only one case was a fishery's most intensely targeted location closed to fishing. Because of the discriminating manner with which fishers target habitats, if simple percentage targets are used for planning, sanctuary location can be adjusted to avoid existing fishing effort. According to modeled outcomes, the implication of this may be diminished reserve effectiveness. To address this, reserve area should be implicitly linked to subducted fishing effort when promoting or modeling MPAs.     

Fishing the line near marine reserves in single and multispecies fisheries.

Throughout the world "fishing the line" is a frequent harvesting tactic in communities where no-take marine reserves are designated. This practice of concentrating fishing effort at the boundary of a marine reserve is predicated upon the principle of spillover, the net export of stock from the marine reserve to the surrounding unprotected waters. We explore the consequences and optimality of fishing the line using a spatially explicit theoretical model. We show that fishing the line: (1) is part of the optimal effort distribution near no-take marine reserves with mobile species regardless of the cooperation level among harvesters; (2) has a significant impact on the spatial patterns of catch per unit effort (CPUE) and fish density both within and outside of the reserve; and (3) can enhance total population size and catch simultaneously under a limited set of conditions for overexploited populations. Additionally, we explore the consequences of basing the spatial distribution of fishing effort for a multispecies fishery upon the optimality of the most mobile species that exhibits the greatest spillover. Our results show that the intensity of effort allocated to fishing the line should instead be based upon more intermediate rates of mobility within the targeted community. We conclude with a comparison between model predictions and empirical findings from a density gradient study of two important game fish in the vicinity of a no-take marine-life refuge on Santa Catalina Island, California (USA). These results reveal the need for empirical studies to account for harvester behavior and suggest that the implications of spatial discontinuities such as fishing the line should be incorporated into marine-reserve design.     

A general bioeconomic simulation model for annual-crop marine fisheries

A generalized bioeconomic simulation model of annual-crop marine fisheries is described and its use in marine fisheries management is demonstrated. The biological submodel represents the recruitment of new organisms into the fishery, the movement of organisms from one fishing area to another and from one depth to another, the growth of organisms and the mortality of organisms resulting both from natural causes and from fishing. The economic submodel represents the fishing effort exerted on each resource species, the monetary costs of fishing, the value of the harvest and the rent (or excess profits) to the fishery.Basic dynamics of the model results from changes in the number of organisms in the fishery over time, which can be summarized as a set of difference equations of the general form ΔN/Δt = R + I ? E ? M ? F where ΔN/Δt is the net change in number of organisms in the fishery over time, R is recruitment, I is immigration, E is emigration, M is natural mortality and F is fishing mortality. R is a driving variable, whereas I, E, M and F are functions of the state of the system at any given point in time. The model can be run in a deterministic or stochastic mode. Values for parameters affecting rates of recruitment, movement, growth, natural mortality and fishing mortality can be selected from uniform, triangular or normal distributions.Use of the model within a fisheries-management framework is demonstrated by evaluating several management alternatives for the pink shrimp (Penaeus duorarum) fishery on the Tortugas grounds in the Gulf of Mexico. Steps involved in use of the model, including parameterization, validation, sensitivity analysis and stochastic simulations of management policies, are explained.     

Striking a Balance between Biodiversity Conservation and Socioeconomic Viability in the Design of Marine Protected Areas

Abstract: The establishment of marine protected areas is often viewed as a conflict between conservation and fishing. We considered consumptive and nonconsumptive interests of multiple stakeholders (i.e., fishers, scuba divers, conservationists, managers, scientists) in the systematic design of a network of marine protected areas along California's central coast in the context of the Marine Life Protection Act Initiative. With advice from managers, administrators, and scientists, a representative group of stakeholders defined biodiversity conservation and socioeconomic goals that accommodated social needs and conserved marine ecosystems, consistent with legal requirements. To satisfy biodiversity goals, we targeted 11 marine habitats across 5 depth zones, areas of high species diversity, and areas containing species of special status. We minimized adverse socioeconomic impacts by minimizing negative effects on fishers. We included fine‐scale fishing data from the recreational and commercial fishing sectors across 24 fisheries. Protected areas designed with consideration of commercial and recreational fisheries reduced potential impact to the fisheries approximately 21% more than protected areas designed without consideration of fishing effort and resulted in a small increase in the total area protected (approximately 3.4%). We incorporated confidential fishing data without revealing the identity of specific fisheries or individual fishing grounds. We sited a portion of the protected areas near land parks, marine laboratories, and scientific monitoring sites to address nonconsumptive socioeconomic goals. Our results show that a stakeholder‐driven design process can use systematic conservation‐planning methods to successfully produce options for network design that satisfy multiple conservation and socioeconomic objectives. Marine protected areas that incorporate multiple stakeholder interests without compromising biodiversity conservation goals are more likely to protect marine ecosystems.     

Collaborative assessment of California spiny lobster population and fishery responses to a marine reserve network

Assessments of the conservation and fisheries effects of marine reserves typically focus on single reserves where sampling occurs over narrow spatiotemporal scales. A strategy for broadening the collection and interpretation of data is collaborative fisheries research (CFR). Here we report results of a CFR program formed in part to test whether reserves at the Santa Barbara Channel Islands, USA, influenced lobster size and trap yield, and whether abundance changes in reserves led to spillover that influenced trap yield and effort distribution near reserve borders. Industry training of scientists allowed us to sample reserves with fishery relevant metrics that we compared with pre-reserve fishing records, a concurrent port sampling program, fishery effort patterns, the local ecological knowledge (LEK) of fishermen, and fishery-independent visual surveys of lobster abundance. After six years of reserve protection, there was a four- to eightfold increase in trap yield, a 5-10% increase in the mean size (carapace length) of legal sized lobsters, and larger size structure of lobsters trapped inside vs. outside of three replicate reserves. Patterns in trap data were corroborated by visual scuba surveys that indicated a four- to sixfold increase in lobster density inside reserves. Population increases within reserves did not lead to increased trap yields or effort concentrations (fishing the line) immediately outside reserve borders. The absence of these catch and effort trends, which are indicative of spillover, may be due to moderate total mortality (Z = 0.59 for legal sized lobsters outside reserves), which was estimated from analysis of growth and length frequency data collected as part of our CFR program. Spillover at the Channel Islands reserves may be occurring but at levels that are insufficient to influence the fishery dynamics that we measured. Future increases in fishing effort (outside reserves) and lobster biomass (inside reserves) are likely and may lead to increased spillover, and CFR provides an ideal platform for continued assessment of fishery-reserve interactions.     

When are no-take zones an economically optimal fishery management strategy?

Discussions on the use of marine reserves (no-take zones) and, more generally, spatial management of fisheries are, for the most part, devoid of analyses that consider the ecological and economic effects simultaneously. To fill this gap, we develop a two-patch ecological-economic model to investigate the effects of spatial management on fishery profits. Because the fishery effects of spatial management depend critically on the nature of the ecological connectivity, our model includes both juvenile and adult movement, with density dependence in settlement differentiating the two types of dispersal. Rather than imposing a reserve on our system and measuring its effect on profits, we ask: "When does setting catch levels to maximize system-wide profits imply that a reserve should be created?" Closing areas to fishing is an economically optimal solution when the value derived from spillover from the reserve outweighs the value of fishing in the patch. The condition, while simple to state in summary form, is complex to interpret because it depends on the settlement success of the dispersing organisms, the nature of the costs of the fishing, the economic and ecological heterogeneity of the system, the discount rate, and growth characteristics of the fish population. The condition is more likely to be satisfied when the closed area is a net exporter of biomass and has higher costs of fishing, and for fish populations with density-independent settlement ("adult movement") than with density-dependent settlement ("larval dispersal"). Rather surprisingly, there are circumstances whereby closing low biological productivity areas, and even sometimes low cost areas to fish, can result in greater fishing profits than when both areas are open to fishing.     

Comanagement Practices Enhance Fisheries in Marine Protected Areas

Abstract:  Fishing activities worldwide have dramatically affected marine fish stocks and ecosystems. Marine protected areas (MPAs) with no-take zones may enhance fisheries, but empirical evidence of this is scant. We conducted a 4-year survey of fish catches around and within an MPA that was previously fully closed to fishing and then partially reopened under regulated comanaged fishing. In collaboration with the fishers and the MPA authority, we set the fishing effort and selected the gear to limit fishing impact on key fish predators, juvenile fish stage, and benthic communities and habitats. Within an adaptive comanagement framework, fishers agreed to reduce fishing effort if symptoms of overfishing were detected. We analyzed the temporal trends of catch per unit of effort (CPUE) of the whole species assemblages and CPUE of the four most valuable and frequent species observed inside the opened buffer zone and outside the MPA investigated. After the comanaged opening, CPUE first declined and then stabilized at levels more than twice that of catches obtained outside the MPA. Our results suggest that working closely with fishers can result in greater fisheries catches. Partial protection of coastal areas together with adaptive comanagement involving fishers, scientists, and managers can effectively achieve conservation and fishery management goals and benefit fishing communities and alleviate overfishing.     

Management and exploitation dynamics of small-scale fisheries in the Bay of Biafra: An integrative analysis of purse seine fishing activity

For many decades, fisheries research has focused on stock assessment and the impact of the fishery effort on resources. Although this knowledge remains necessary, a more integrated analysis of the joint dynamics of resource and operational activities is needed to provide more useful advice for the management of fishery systems. Since 1994 a new approach to fishery science has been carried out for Cameroonian small-scale fisheries, the aim being the incorporation into fishery science of research on fishery management, fishing processes and fishermen's behaviour. This paper presents a more systemic data approach which combines biological parameters and operational factor analyses with the goal of sustainable development. From two years of data collected on the purse seine fishing units operating in the Bay of Biafra (2002 and 2003), a set of three correspondence analyses is applied: (1) to the length frequency distribution of Ethmalosa fimbriata, (2) to the number of visits per fishing ground, and (3) to the species appearance frequency in the landings. These three analyses were plotted per month-year period. The results are presented in the form of an annual exploitation cycle, linking fishing grounds, the main species caught, and corresponding fishing period.     

Incorporation of Spatial and Economic Analyses of Human‐Use Data in the Design of Marine Protected Areas

Abstract: Social, economic, and ecological criteria contribute to the successful design, implementation, and management of marine protected areas (MPAs). In the context of California's Marine Life Protection Act Initiative, we developed a set of methods for collecting, compiling, and analyzing data about the spatial extent and relative economic importance of commercial and recreational fishing. We interviewed 174 commercial fishers who represented the major fisheries in the initiative's north‐central coast region, which extends from Point Arena south to Pigeon Point. These fishers provided data that we used to map the extent of each of the fishing grounds, to weight the relative importance of areas within the grounds, to characterize the operating costs of each fishery, and to analyze the potential economic losses associated with proposed marine protected areas. A regional stakeholder group used the maps and impact analyses in conjunction with other data sets to iteratively identify economic and ecological trade‐offs in designations of different areas as MPAs at regional, port, and fishery extents. Their final proposed MPA network designated 20% of state waters as MPAs. Potential net economic loss ranged from 1.7% to 14.2% in the first round of network design and totaled 6.3% in the final round of design. This process is a case study in the application of spatial analysis to validate and integrate local stakeholder knowledge in marine planning.     

Modeling spatial patterns in fisheries bycatch: improving bycatch maps to aid fisheries management.

Fisheries bycatch, or incidental take, of large vertebrates such as sea turtles, seabirds, and marine mammals, is a pressing conservation and fisheries management issue. Identifying spatial patterns of bycatch is an important element in managing and mitigating bycatch occurrences. Because bycatch of these taxa involves rare events and fishing effort is highly variable in space and time, maps of raw bycatch rates (the ratio of bycatch to fishing effort) can be misleading. Here we show how mapping bycatch can be enhanced through the use of Bayesian hierarchical spatial models. We compare model-based estimates of bycatch rates to raw rates. The model-based estimates were more precise and fit the data well. Using these results, we demonstrate the utility of this approach for providing information to managers on bycatch probabilities and cross-taxa bycatch comparisons. To illustrate this approach, we present an analysis of bycatch data from the U.S. gill net fishery for groundfish in the northwest Atlantic. The goals of this analysis are to produce more reliable estimates of bycatch rates, assess similarity of spatial patterns between taxa, and identify areas of elevated risk of bycatch.     

Sensitivity of Marine-Reserve Design to the Spatial Resolution of Socioeconomic Data

Abstract:  Socioeconomic considerations should have an important place in reserve design. Systematic reserve-selection tools allow simultaneous optimization for ecological objectives while minimizing costs but are seldom used to incorporate socioeconomic costs in the reserve-design process. The sensitivity of this process to biodiversity data resolution has been studied widely but the issue of socioeconomic data resolution has not previously been considered. We therefore designed marine reserves for biodiversity conservation with the constraint of minimizing commercial fishing revenue losses and investigated how economic data resolution affected the results. Incorporating coarse-resolution economic data from official statistics generated reserves that were only marginally less costly to the fishery than those designed with no attempt to minimize economic impacts. An intensive survey yielded fine-resolution data that, when incorporated in the design process, substantially reduced predicted fishery losses. Such an approach could help minimize fisher displacement because the least profitable grounds are selected for the reserve. Other work has shown that low-resolution biodiversity data can lead to underestimation of the conservation value of some sites, and a risk of overlooking the most valuable areas, and we have similarly shown that low-resolution economic data can cause underestimation of the profitability of some sites and a risk of inadvertently including these in the reserve. Detailed socioeconomic data are therefore an essential input for the design of cost-effective reserve networks.     

Commercial fisheries under price uncertainty

The deterministic models applied in economics of fisheries are extended to comprise price uncertainty and risk aversion among the fishing units. It is proved that in the open-access fishery both the total fishing effort and the number of fishing units are reduced as the variance of the price increases; that the total fishing effort may be smaller in the open-access fishery than in the optimal fishery at a high variance; that only a fixed producer price system can create a first-best optimum, and that a tax on revenue is more efficient than both fishing unit quotas or tax on catch.     

Fishing‐gear restrictions and biomass gains for coral reef fishes in marine protected areas

Considerable empirical evidence supports recovery of reef fish populations with fishery closures. In countries where full exclusion of people from fishing may be perceived as inequitable, fishing‐gear restrictions on nonselective and destructive gears may offer socially relevant management alternatives to build recovery of fish biomass. Even so, few researchers have statistically compared the responses of tropical reef fisheries to alternative management strategies. We tested for the effects of fishery closures and fishing gear restrictions on tropical reef fish biomass at the community and family level. We conducted 1,396 underwater surveys at 617 unique sites across a spatial hierarchy within 22 global marine ecoregions that represented 5 realms. We compared total biomass across local fish assemblages and among 20 families of reef fishes inside marine protected areas (MPAs) with different fishing restrictions: no‐take, hook‐and‐line fishing only, several fishing gears allowed, and sites open to all fishing gears. We included a further category representing remote sites, where fishing pressure is low. As expected, full fishery closures, (i.e., no‐take zones) most benefited community‐ and family‐level fish biomass in comparison with restrictions on fishing gears and openly fished sites. Although biomass responses to fishery closures were highly variable across families, some fishery targets (e.g., Carcharhinidae and Lutjanidae) responded positively to multiple restrictions on fishing gears (i.e., where gears other than hook and line were not permitted). Remoteness also positively affected the response of community‐level fish biomass and many fish families. Our findings provide strong support for the role of fishing restrictions in building recovery of fish biomass and indicate important interactions among fishing‐gear types that affect biomass of a diverse set of reef fish families.     

Modeling spawning dates of Hucho taimen in Mongolia to establish fishery management zones.

The ecological impacts of recreational fisheries are of growing concern and pose a number of unique management challenges. Here we report on our efforts to provide guidance for managing a recreational fishery for taimen, the giant Eurasian trout (Hucho taimen) in Mongolia. This species has declined dramatically across its range of Siberia and Central Asia, and is currently listed as endangered in Mongolia. Strong populations persist in remote regions of Mongolia because of limited anthropogenic impacts and harvest, though interest in the fishery is expanding rapidly. Current fishing regulations list the spring "opening date" for taimen fishing as 15 June, although regulations have not been consistently enforced, partially because taimen spawn much earlier than 15 June in much of the country. Through a combination of statistical models, climate data, knowledge of taimen biology, and geographic information systems (GIS), we model taimen spawning dates for potential habitat in Mongolia. A parametric bootstrap procedure was used to simulate variability in spawning date derived from inter-annual climate variability and model error, from which we estimated the date in which taimen spawning is predicted to occur with 90% confidence. We recommend the designation of three fisheries management zones, with corresponding opening dates of 20 May, 1 June, and 15 June. Our fishery opening date recommendations are less restrictive than existing regulations. Provided there is little or no catch-and-release fishing mortality, this approach serves both environmental and human needs by protecting taimen during the reproductive period, while still allowing a post-spawning catch-and-release fishery that benefits local economies and generates revenue (through fishing concession fees) for local conservation efforts.     

Effects of sex change on the implications of marine reserves for fisheries

Marine reserves have become widely used in biodiversity conservation and are increasingly proposed as fisheries management tools. Previous modeling studies have found that reserves may increase or decrease yields, depending on local environmental conditions and on the specific life-history traits of the fishery species. Sex-changing (female-to-male) fish are targets of some of the most important commercial and recreational fisheries in the world. The potential for disproportionate removal of the larger, older sex of such species requires new theory to facilitate our understanding of how reserves will affect the yields of surrounding fisheries, relative to fishes with separate sexes. We investigated this question by modeling the effects of marine reserves on a non-sex-changing and a sex-changing population. We used demographic parameter estimates for the common coral trout as a baseline, and we conducted extensive sensitivity analyses to determine how sustainable yields of sex-changing species are likely to be affected by reserves across a broad range of life-history parameters. Our findings indicate that fisheries for sex-changing species are unlikely to receive the same yield-enhancing benefit that non-sex-changing fisheries enjoy from marine reserves, and that often reserves tend to reduce sustainable yields for a given overall population size. Specifically, the increased egg production and high fertilization success within reserves is more than offset by the reduced egg production and fertilization success in the fished areas, relative to a system in which fishing mortality is distributed more evenly over the entire system. A key reason for this appears to be that fertilization success is reduced, on average, when males are unevenly distributed among subpopulations, as is the case when reserves are present. These findings suggests that, for sex-changing populations, reserves are more suited to rebuilding overfished populations and sustaining fishery viability, rather than enhancing fishery yields. These results are robust over a range of sex-change regimes, stock-recruitment relationships, adult mortality rates, individual growth strategies, and fertilization-success functions. Our findings highlight the importance of considering the different contributions of males and females to population growth and fishery yields when evaluating the efficacy of marine reserves for enhancement of fished species.     

Effects of Fisheries Closures and Gear Restrictions on Fishing Income in a Kenyan Coral Reef

Abstract: The adoption of fisheries closures and gear restrictions in the conservation of coral reefs may be limited by poor understanding of the economic profitability of competing economic uses of marine resources. Over the past 12 years, I evaluated the effects of gear regulation and fisheries closures on per person and per area incomes from fishing in coral reefs of Kenya. In two of my study areas, the use of small‐meshed beach seines was stopped after 6 years; one of these areas was next to a fishery closure. In my third study area, fishing was unregulated. Fishing yields on per capita daily wet weight basis were 20% higher after seine‐net fishing was stopped. The per person daily fishing income adjacent to the closed areas was 14 and 22% higher than the fishing income at areas with only gear restrictions before and after the seine‐net restriction, respectively. Incomes differed because larger fish were captured next to the closed area and the price per weight (kilograms) increased as fish size increased and because catches adjacent to the closure contained fish species of higher market value. Per capita incomes were 41 and 135% higher for those who fished in gear‐restricted areas and near‐closed areas, respectively, compared with those who fished areas with no restrictions. On a per unit area basis (square kilometers), differences in fishing income among the three areas were not large because fishing effort increased as the number of restrictions decreased. Changes in catch were, however, larger and often in the opposite direction expected from changes in effort alone. For example, effort declined 21% but nominal profits per square kilometer (not accounting for inflation) increased 29% near the area with gear restrictions. Gear restrictions also reduced the cost of fishing and increased the proportion of self‐employed fishers.     

Aquaculture and the displacement of fisheries captures

In modern aquaculture, animal-production technology is used to increase aquatic food sources. Such controlled rearing of seafood can, in principle, shift the pressure off wild stocks and aquatic ecosystems by reducing fishing activities, which may advance marine conservation goals. We examined resource displacement—the reduced consumption of a resource due to its replacement with a more environmentally benign substitute—in fisheries. We employed panel regression techniques in an analysis of time-series data from 1970 through 2014 to assess the extent to which aquaculture production displaced fisheries captures for all nations for which data were available. We estimated 9 models to assess whether aquaculture production suppresses captures once other factors related to demand have been controlled for. Only 1 model predicted significant suppression of fisheries captures associated with aquaculture systems within nations over time. These results suggest that global aquaculture production does not substantially displace fisheries capture; instead, aquaculture production largely supplements fisheries capture.     

Marine reserves,fisheries ban,and 20 years of positive change in a coral reef ecosystem

By 2004, Belize was exhibiting classic fishing down of the food web. Groupers (Serranidae) and snappers (Lutjanidae) were scarce and fisheries turned to parrotfishes (Scarinae), leading to a 41% decline in their biomass. Several policies were enacted in 2009–2010, including a moratorium on fishing parrotfish and a new marine park with no-take areas. Using a 20-year time series on reef fish and benthos, we evaluated the impact of these policies approximately 10 years after their implementation. Establishment of the Southwater Caye Marine Reserve led to a recovery of snapper at 2 out of 3 sites, but there was no evidence of recovery outside the reserve. Snapper populations in an older reserve continued to increase, implying that at least 9 years is required for their recovery. Despite concerns over the feasibility of banning parrotfish harvest once it has become a dominant fin fishery, parrotfishes returned and exceeded biomass levels prior to the fishery. The majority of these changes involved an increase in parrotfish density; species composition and adult body size generally exhibited little change. Recovery occurred equally well in reserves and areas open to other forms of fishing, implying strong compliance. Temporal trends in parrotfish grazing intensity were strongly negatively associated with the cover of macroalgae, which by 2018 had fallen to the lowest levels observed since measurements began in 1998. Coral populations remained resilient and continued to exhibit periods of net recovery after disturbance. We found that a moratorium on parrotfish harvesting is feasible and appears to help constrain macroalgae, which can otherwise impede coral resilience.     

Malthusian overfishing and efforts to overcome it on Kenyan coral reefs

This study examined trends along a gradient of fishing intensity in an artisanal coral reef fishery over a 10-year period along 75 km of Kenya's most populated coastline. As predicted by Malthusian scenarios, catch per unit effort (CPUE), mean trophic level, the functional diversity of fished taxa, and the diversity of gear declined, while total annual catch and catch variability increased along the fishing pressure gradient. The fishery was able to sustain high (approximately 16 Mg x km(-2) x yr(-1)) but variable yields at high fishing pressure due to the dominance of a few productive herbivorous fish species in the catch. The effect of two separate management strategies to overcome this Malthusian pattern was investigated: fisheries area closure and elimination of the dominant and most "competitive" gear. We found that sites within 5 km of the enforced closure showed significantly lower total catch and CPUE, but increased yield stability and trophic level of catch than predicted by regression models normalized for fishing effort. Sites that had excluded illegal beach seine use through active gear management exhibited increased total catch and CPUE. There was a strong interaction between closure and gear management, which indicates that, for closures to be effective at increasing catch, there must be simultaneous efforts at gear management around the periphery of the closures. We propose that Malthusian effects are responsible for the variation in gear and catch and that active management through reduced effort and reductions in the most competitive gear have the greatest potential to increase the functional and trophic diversity and per-person productivity.